Eolid Mollusca from Jamaica, with Descriptions of Two New Genera and Three New Species

BULLETIN OF MARINE SCIENCE OF THE GULF AND CARIBBEAN

VOLUME 14 1964 NUMBER 1

EOLID MOLLUSCA FROM JAMAICA, WITH DESCRIPTIONS OF TWO NEW GENERA AND THREE NEW SPECIES

MALCOLM EDMUNDS Department of Zoology, University of Ghana

ABSTRACT Thirteen species of eolid nudibranchs were collected between October 1961 and March 1962 from mangrove roots at Port Royal, Jamaica. An additional small collection was made in Biscayne Bay, Florida, near the Institute of Marine Science, University of Miami, on April 2, 1962. Two new genera are established, Selva (Cuthonidae) and Palisa (Facelinidae), as well as three new species, Selva rubra, Palisa papillata and Godiva rubrolineata. The classification of the subfamily Facalaninae is discussed. Of the 13 species of eolids known from Jamaica, 10 occur also at Miami and 8 near Sao Paulo, Brazil. Four instances of eolids being parasitized by pycnogonids are described.

INTRODUCTION The material described in this paper was collected whilst the author was working in the Department of Zoology of the University of the West Indies, Mona, Jamaica. All of the Jamaican animals were found at Port Royal where the Department of Zoology has a small marine laboratory. One individual each of Berghia coerulescens and Spurilla neapolitana, and small numbers each of Catriona tina, Favorinus auritulus and Palisa papillata were found in tanks at Port Royal or on hydro ids growing nearby. The rest of the material was collected from a narrow channel in the mangroves near Port Royal, at Goodbody's station A (Goodbody, 1961). The water in this channel varies in depth from 3-4 inches to 3-4 feet, and the stems and roots of the mangroves are covered with a rich growth of hydroids and anemones on which the eolids feed. Collections were made at irregular intervals between October 1961 and March 1962. A further small collection was made from hydroids growing on dock pilings on Key Biscayne, close to the Marine Laboratory of the University of Miami, Florida, on April 2, 1962. The nudibranchs from ·the Caribbean have been studied recently by Marcus and Marcus (1960), but prior to this, the only reasonably com- prehensive study is that of Engel (1925, 1927). Since there is a close resemblance between the opisthobranchs from Florida and from the region of Sao Paulo, Brazil (Marcus and Marcus, 1960), the papers by Marcus 2 Bulletin of Marine Science of the Gulf and Caribbean [14(1) (1955, 1957, 1958) on Brazilian opisthobranchs have also proved to be valuable. The classification used in the present paper is based upon that of Odhner (1939), modified by Macnae (1954) and Marcus (1957, 1958). In this paper, 13 species of eolids are discussed, including two new genera, Palisa and Selva, and three new species, Pa/isa papillata, Selva rubra and Godiva rubrolineata. The holotypes of these new species are deposited in the British Museum (Natural History), London. EXPLANATION OF LETTERING ace-accessory gland m-masticatory border amp-ampulla ovi-oviduct an-anus ovo-ovotestes ant.l-anterior liver p-penis cn-cnidosac post. I-posterior liver d-dorsal indentation pr-prostate I.g.a-female genital atrium p.t-pre-radular tooth j.g.m-female gland mass r-rectum g-gonopore r.p--renal pore g.a-genital atrium s-stomach h-heart sp.ov-spermoviduct h.d-hermaphrodite duct sp.th-sperrnatheca hi-hinge st-stylet k-kidney v-vagina I-liver v.d-vas deferens Order NUDIBRANCHIA Suborder EOLIDACEA Tribe ACLEIOPROCTA Family Eubranchidae Capellinia canida Marcus Occurrence.- Two animals were found in January and February 1962 at Port Royal. Further distribution.-At two localities in Brazil and at two localities in Florida (Marcus, 1958; Marcus and Marcus, 1960). Discussion.-The two animals were 3.0 and 2.5 mm long when alive, and closely resemble the specimens described by Marcus (1958) and by Marcus and Marcus (1960). There is considerable color variation in this species, especially in the amount of white and of brown. The radula and the reproductive organs were examined to confirm the identification. Family Cuthonidae Catrio1Ulmaua Marcus and Marcus Figs. 1, A-B; 2, B-C; 3, A Occurrence.-Forty-two specimens were found between October 1961 and March 1962. 1964] Edmunds: Eolid Mollusca 3 Further distribution.-Miami, Florida (Marcus and Marcus, 1960). Description.-The largest animal was 12 mm long when alive, but the usual length is 5 to 8 mm. The original description of C. maua is based upon observations of two preserved specimens (Marcus and Marcus, 1960), so many of the details which follow are given for the first time. The living animal has the typical rounded foot and smooth rhinophores of the genus Catriona. In an animal 9 mm long, the cerata reach 3 mm, the oral tentacles 2 mm, and the rhinophores 3 mm in length. There is a white stippling on the distal halves of both rhinophores and oral tentacles, and there are further white markings on the head and back (Fig. 1, A). Occasionally, the white markings are tinged with orange. There is a red streak up the basal half of the rhinophores on the posterior side. The cerata have two white bands toward the tip (Fig. 1, B), and there is often a faint suffusion of orange to the otherwise colourless cera tal epidermis. Creamy-white unicellular epidermal glands are present in the white band nearest the tip of the ceras. The liver in the cerata of fresh animals is pale red with cream and dark red blotches, but after a few days in captivity the red changes to pale brown. The liver ducts in the body may be slightly pinkish, and the oesophagus is bright red. The

FIGURE 1. A, Catriona maua Marcus & Marcus: dorsal view of living animal; white markings shown by fine stippling, red by coarse stippling.-B, Ceras of C. maua; white bands are stippled; the liver is pale red with dark red 3;nd cream blotches.-C, Catriona perea Marcus: dorsal view of living animal; white markings finely stippled, brown and blackish markings coarsely stippled. -D, Ceras of C. perea; white markings stippled; liver is yellow-brown mostly obscured by black. 4 Bulletin of Marine Science of the Gulf and Caribbean [14(1) general body colour is silvery-grey, sometimes faintly suffused with orange, though never as much as in C. aurantia. The cerata stand on cushions, as described by Marcus and Marcus (1960), and in a large (9 mm) animal have the following distribution: Anterior liver Posterior liver Right side 6,5,5 3,3,3,1,1,1 Left side 6,5,5 4,3,1,3,1,1 The anus lies just above the first ceras of the posterior liver, and the genital openings are either just in front of or just behind the base of the first row of cerata of the anterior liver. The jaws resemble those described by Marcus and Marcus (1960), but there is no black pigment in any of four animals examined. The radula has up to 100 teeth (Fig. 2,B). Since the pre-radular tooth is retained in this species (Fig. 2,C), the number of teeth indicates age and is not of taxonomic significance.

C 20" .

~~~ p.l

FIGURE 2. A, Catriona perea Marcus: 17th tooth from specimen with 26 radular teeth.-B, Catriona maua Marcus & Marcus: 87th tooth from specimen with 95 radular teeth.-C, Catriona maua: first five radular teeth, induding the pre-radular tooth.

The genital system is similar to that of C. perea. In one animal there are four testis sacs, each with six or more ovaries opening into them. After leaving the 0.2 mm-Iong ampulla, the spermoviduct divides into oviduct and vas deferens (Fig. 3,A). Most of the vas deferens is prostatic, except for a short length near the penis. There is an accessory male gland, but this differs from the accessory male gland of C. perea in that the cells are columnar and closely resemble those of the prostate gland. The spermatheca is a pouch 80p. in diameter, opening into the female atrium close to the oviduct. The penial stylet is 3'O,ulong. Catriona perea Marcus Figs. 1, C-D; 2, A; 3, B Occurrence.-Thirty-one specimens were found at Port Royal in Novem- ber and Decemer 1961. 1964] Edmunds: Eolid Mollusca 5 Further distribution. - Brazil (Marcus, 1958). Description. - The largest animal, 15 mm long when alive, was decaying slightly and was covered with epizootic protozoa. The living animal is exactly as described by Marcus (1958), but the oral tentacles and rhinophores (Fig. 1, C of this paper) are stouter than in his figure. The most important features are the orange pigment in the head region (not indicated in Fig. 1, C), the blotch of brown or black pigment in front of the rhinophores, and the blackish liver ducts which are clearly visible through the epidermis. The foot is rounded as in other species of Catriona. The liver in the cerata is basically yellow-brown, but this is largely obscured by black tubercles (Fig. 1, D). There are two bands of white on the cerata, and, as in C. maua, there are white glands in the band nearest the tip.

FIGURE 3. A, Catriona maua Marcus & Marcus.-B, Catriona perea Marcus. Diagrams of reproductive system. 6 Bulletin of Marine Science of the Gulf and Caribbean [14(1) The cerata are distributed as described by Marcus (1958), and in one animal 10 mm long had the following numbers per row: Anterior liver Posterior liver Right side 7,8 8,3,5,3,3,1,1,1 Left side 7,8 6,4,4,3,2,1,1,1 glvmg a total of 77. An animal 11.5 mm long had 98 cerata, whilst an animal 7 mm long had only 38. The anus is closely in front of the first row of the posterior liver. The jaws and radula are as described by Marcus (1958). In three animals which were examined there are 16, 22, and 26 teeth in the radula. The largest and the smallest teeth from each radula have the following breadths: 108 and 87p" 78 and 42.5p. and 85 and 58p, respec- tively. There are from 6 to 10 lateral dentic1es to the cusp of each radular tooth, and the longest of these denticles is considerably longer than the cusp. The dentic1es often are of irregular length, as described by Marcus, and in this respect Fig. 2A is atypical. The genital system of the present material is very similar to that of Brazilian animals described by Marcus. The genital openings lie just in front of the first row of cerata on the right side. There are up to 10 pairs of testis follicles, into each of which open several ovaries. In one large animal, the vagina is a pouch which contains sperm, and the spermathecal duct opens near it (Fig. 3, B).

Catriona tina Marcus Figs. 4; 5, A Occurrence.-Sixty-eight specimens were found at Port Royal between November 1961 and March 1962. Further distribution.-Brazil and Florida (Marcus, 1957; Marcus and Marcus, 1960). Description.- The living animal is up to 4.5 mm long, with a rounded or slightly notched foot. The oral tentacles are rather less than 1 mm long; the rhinophores are smooth, and up to 1 mm in length. The color of the animal is very variable. The entire dorsal surface may be covered with white dots, or these may be confined to an irregular median line, or they may be thinly and irregularly scattered. Usually there is a band of white near the tips of the oral tentacles and the rhinophores. There is often some orange pigment in front of the heart, by the first ceras row, at the base of the rhinophores, and sometimes at the base of the oral tentacles; but the orange may be pale and in some cases is completely lacking. Sometimes the rhinophores are pale yellow at the base and lack orange. The cerata have one white ring near the tip 1964] Edmunds: EoUd Mollusca 7 which contains white epidermal glands, but the disposition of white over the rest of the ceratal surface is very variable. The liver is yellow- brown with scattered darker patches, especially at the base of the ceras. There are up to 11 cerata in five rows on either side of the body. The anterior livers give off two rows on either side, each with 2 or 3 cerata per row. In the posterior liver the maximum number of cerata per row is 3. The jaws and radula resemble those described by Marcus (1957) and Marcus and Marcus (1960). The radula of one animal has 39 teeth with two more in process of formation. Teeth from two animals are illustrated in Fig. 4.

20 ~. B A FIGURE 4. Catriona tina Marcus: radular teeth. A, Last tooth of an animal with 39 teeth.-B, 38th tooth of an animal with 39 teeth.

The reproductive system is as described by Marcus (1957). In one animal that was sectioned there are five testis sacs, into which open a number of female follicles. Important taxonomic features are the short, broad prostate (150,.. long by 90,.. broad), the accessory male gland with columnar epithelium 20fL high, and the curved penial stylet up to 80,.. long (Fig. 5, A).

Selva, n. gen. Diagnosis.-Eolidacea with the anus in the c1eioproct position; with two unbranched rows of cerata to the anterior liver; rhinophores smooth; foot rounded anteriorly; radula uniseriate, the central cusp projecting beyond the lateral denticles; with the spermatheca opening into the female atrium and separated from the spermoviduct by the whole of the female gland mass; with the penis armed; and with an accessory male gland. Selva can be distinguished from all other acleioprocts by the cIeioproct anus and by the combination of characters given here. It belongs to the family Cuthonidae and is closely related to Tergipes, Catriona and Cuthona. Type-species.-Selva rubra, n. sp. 8 Bulletin of Marine Science of the Gulf and Caribbean [14(1)

FIGURE 5. A, Catriona tina Marcus.-B, Selva rubra, n. sp. Diagrams of reproductive system.

Selva rubra, n. sp. Figs. 5, B; 6; 7 Occurrence.- Three specimens were found between January and March 1962 in the mangroves at Port Royal. The holotype, fixed in Heidenhain's "Susa," is in the British Museum (N. H.), no. 1963115 W. Description.-The living animal is up to 4.5 mm long including the short tail, which does not extend far behind the last row of cerata. The general body color is silvery gray. The rhinophores are smooth, up to 1 mm long, the distal half speckled with white. The oral tentacles are up to 0.75 mm long, with the distal three-quarters white. White coloring is also present on the forehead, behind the eyes, on the sides of the head, as a broken band along the flanks, and dorsally on the tail (Fig. 6, A). Orange pigment is present in a streak from the basal 1964] Edmunds: Eolid Mollusca 9

1.0 1mI

0.5 nun c

FIGURE6. Selva rubra, n. sp. A, Dorsal view of living animal: white markings finely stippled, orange coarsely stippled.-B, Ventral view of head, showing shape of foot.-C, Right side with three anterior cerata removed: stippling as in A.-D, Ceras: dark stippling represents crimson, light stippling brown, and circles are cream spots. one-quarter of the oral tentacles up the forehead, and up the front half of the rhinophores. There is another orange streak running forward from the first group of cerata and up the posterior two-thirds of the rhinophores; and there is a complete ring of orange about one-third of the way up the rhinophores (Fig. 6, A and C). The liver is clearly visible through the transparent body. It is colored crimson at the anterior and posterior ends and brown in the middle. There are two rows of cerata to the anterior liver and three to the posterior liver, but there is a projection of the liver in the tail of the eolid behind the last row of cerata, suggesting that more rows may occur in larger animals. The largest animal has the following numbers of cerata per row beginning at the anterior end: 2, 2, 2, 1, 1 on the right and 2,3,2,1, 1 on the left. The cerata are up to 1 mm long with the cnidosac up to 0.25 mm in diameter (Fig. 6, D). They are broadest in the cnidosac region, as are the cerata of Tergipes despectus (Alder and Hancock, 1845-1855). The cnidosac is white, and there is sometimes a white blotch in the epidermis above it. The liver in the cerata is crimson just below the cnidosac, merging into a brownish color with cream flecks below. 10 Bulletin of Marine Science of the Gulf and Caribbean [14(1) The genital aperture is just below and anterior to the first cerata on the right side of the body. The anus is immediately posterior to the first rOw of cerata of the posterior liver on the right side, that is, in the cleioproctic position. The renal pore is in the interhepatic space, just in front of the first cerata of the posterior liver. The foot is rounded anteriorly and is not notched in life (Fig. 6, B), although it may have a very shallow notch when fixed. The gonads are visible through the transparent body of the animal in life and are cream colored. The jaws of the largest animal are 0.42 mm in length with a strong hinge (Fig. 7, A). The masticatory border is roughened, but is not denticulate. The radula of the same animal has 31 teeth excluding a "ghost" tooth in process of formation, whilst a second animal of 4 mm overall length has 24 teeth. Each tooth has a strong median cusp with from 6 to 9 long, finely pointed denticles on either side of it. The cusp is considerably longer than the flanking denticles. The larger and more recent teeth do not differ markedly from the older and smaller teeth (Fig. 7, B and C).

20 ..;.

FIGURE 7. Selva rubra, n. sp. A, Jaw.-B, 29th radular tooth from animal with 31 teeth.-C, 8th tooth of same animal.

The cerata contain numerous special cells ("cellules spcciales" according to Hecht, 1896), similar to those found in certain other eolids. The kidney is similar to that of Tergipes despectus, as described and figured by Marcus (1957) but there are two openings from the kidney tube to the pericardium, both on the right side and both with a bunch of long cilia. The reproductive system is similar to that found in species of Catriona and Tergipes (Fig. 5, B) There are two testis sacs, into each of which open a number of ovarian pouches. From these two sacs, the 8wdiameter hermaphrodite duct runs forward to enter the ampulla at its anterior end. The ampulla is not a swelling of the hermaphrodite duct as it is in Catriona perca, but is a pouch 0.2 mm long. 1964] Edmunds: Eolid Mollusca 11 It is probable that eggs can pass down to the oviduct without entering the ampulla at all. Shortly after leaving the ampulla, the spermoviduct divides into the vas deferens and the oviduct. The vas deferens is prostatic for most of its length, and in the penis it receives the duct from an accessory male gland. This accessory gland has the same histological structure as the accessory male gland of Catriona perea. The penis bears at its tip a hollow stylet, 10JL in diameter at its base, 5JL at its tip, and 181-' long in the specimen 4.5 mm long. Eggs pass down the oviduct into the female gland mass where they presumably receive mucous and albumen secretions. They then enter the female atrium and pass out through the common genital opening. The spermatheca is a pouch of 1301-' diameter which opens into the female atrium. Copulation was not observed, but the presence of sperm in the spermatheca indicates that this species is mature at a length of 4.5 mm. Discussion.-Following the system of classification of Odhner (1939) the Eolidacea are divided into three groups according to the position of the anus; the Pleuroprocta with the anus lateral to the cerata, the Acleioprocta with the anus in the interhepatic space, and the Cleioprocta with the anus between the cerata of the posterior liver. According to this system, Selva is a c1eioproct. However, the combination of an armed penis, cuspidate radular teeth, smooth rhinophores, rounded foot, and two short rOws of cerata to the anterior liver distinguish it from all cleioproct genera. It differs from the Aeolidiidae in the cuspidate radular teeth, and from the Facelinidae and the Favorinidae in the position of the spermatheca. In these two families, there is a short duct from the spermatheca which enters the oviduct close to the point at which the spermoviduct divides into the vas deferens and the oviduct (Odhner, ]939; Macnae, 1954). In the cuthonids Catriona perea and TerRipes despectus, the spermathecal duct enters the female genital atrium, and is separated from the spermoviduct by the whole of the female gland mass. This is identical with the situation found in Selva. Thus, Selva differs in many ways from c1eioproct genera and families. In all of these features it resembles the condition found in ac1eioprocts, particularly the Cuthonidae, and it is necessary to consider whether it might be an acleioproct which has secondarily acquired the c1eioproct condition. Selva resembles the Cuthonidae and differs from typical cleioprocts in the following features: the position of the spermatheca; the presence of an accessory male gland of similar histological structure to that of Catriona perea; the possession of a hollow penial stylet; the position of the gonopore just anterior to the first row of cerata; the smooth rhino- phares; the rounded foot without a conspicuous notch; the distribution of the cerata of the anterior liver in two short rows; the absence of denticles on the masticatory border of the jaw; and the possession of 12 Bulletin of Marine Science of the Gulf and Caribbean [14(1) "cellules speciaIes" in the cerata. The radula with a projecting central cusp and finely pointed lateral denticles is similar to radulae from both c1eioprocts and acleioprocts, but it debars Selva from the genus Catriona in which the central cusp is not markedly longer than the lateral cusps. The blunt cerata of Selva, with their large cnidosacs, resemble the condition found in Tergipes despectus, but are not described, to my knowledge, from any cleioproct. These features conclusively demon- strate that Selva rubra is a cuthonid which has secondarily acquired a cleioproct anus. Precuthona is another acleioproct in which the most anterior cerata of the posterior liver may arise in front of the anus (Odhner, 1939; Marcus, 1961). In Precuthona each liver branch divides into several irregular rows, one or two of which may lie just anterior to the anus. In Selva there are few cerata per row, the rows are unbranched, and one complete row lies anterior to the anus. Thus the cleioproct condition has evolved at least twice in the acleioproct eolids.

Tribe CLEIOPROCTA Family Facelinidae PaJisa, n. gen. Diagnosis.-Facelinidae with tuberculate rhinophores; radular teeth lacking a prominent cusp, the denticles irregularly serrulate; no accessory male gland; penis unarmed and not containing the prostate gland; spermatheca forming a part of the oviduct. Type-species.-Palisa papillata, n. sp. Remarks.-This diagnosis may require modification when more species are described in the genus, but at present it is impossible to decide which characters should be given specific rank and which are of generic importance. Facelinid genera are in need of a thorough revision, but until the reproductive systems are better known it is difficult to find good generic characters. Facelina agari Smallwood from Bermuda is at present poorly known (Smallwood, 1910), but it may belong to Palisa. F. agari, Moridilla and Noumeaella Risbec have papillae on the rhinophores, but the ornamen- tation of the rhinophores is so variable in the Facelinidae that this need not indicate that they are congeneric. Noumeaella may perhaps be a facelinid, but it has an armed penis (Risbec, 1937, 1953). Palisa papilJata, n. sp. Figs. 8; 9; 10, A Occurrence.- Twenty-four individuals were found between October 1961 and February 1962 at Port Royal. A single specimen was found at Miami in April 1962. The holotype, from Jamaica, is fixed in Heiden- 1964] Edmunds: Eolid Mollusca 13 hain's "Susa" and is deposited in the British Museum (N.H.), no. 1963116 W. Description.- The living animal is up to 16 mm long, with a narrow body and long, pointed tail (Fig. 8, A, C). The foot is notched anteriorly, longitudinally grooved, and has tentacle-like anterior corners which may reach 2 mm in length (Fig. 8, B). The oral tentacles are very mobile and are as much as 5 mm in length. An animal 8 mm in length has oral tentacles 2.5 mm long, and projections 1 mm long at the anterior angles of the foot. The rhinophores are smooth on the anterior face, but the posterior surface is covered with small papillae similar to those found in Berghia coerulescens. An animal 8 mm long has 42 papillae in irregular oblique rows on its rhinophores, which are 0.9 mm long (Fig. 8, E). An animal 3 mm long has only 6 or 7 papillae on each rhinophore. The body is silvery-gray in color, but the whole of the dorsal surface, including the cerata, is covered with a meshwork of fine white lines which may form an almost solid patch on the back of large animals. The viscera are pale buffish, or silvery-grey like the body, but

FIGURE 8. Palisa papillata, n. sp. A, dorsal view of living animaL-B, Ventral view of head showing notched foot.-C, View of right side with cerata removed.-D, Ceras, showing the irregular white lines and the darker spots in the liver.-E, Contracted rhinophore with papillae. 14 Bulletin of Marine Science of the Gulf and Caribbean [14(1) in large specimens are tinged with orange. The liver in the cerata is pale buffish, but there are usually dark brown spots just below the cnidosac, at the base of the ceras, and sometimes in between as well (Fig. 8, D). The cerata may reach 2.5 mm in length in a large animal, and the cnidosac has a very long duct to the cnidopore. The nemato- cysts are not enclosed in packets (or cnidocysts) as is usual in other eolids, but lie freely in the cnidosac. They are all isorhizas 18-24~ long and 11 to 12,u broad, so that they can pass easily down the 12~-wide duct to the cnidopore. The arrangement of cerata is similar to that described for Learchis poica (Marcus and Marcus, 1960). The anterior livers give off 3, 4, or 5 rows of cerata, whilst the posterior liver gives off cerata in simple arches toward the front, and in rows farther back (Fig. 8, C). The numbers of cerata per liver duct and the total numbers of cerata in animals of various sizes are given in Table 1.

TABLE 1 NUMBER OF CERATA PER ROW IN SPECIMENS OF Palisa pap illata Length of Side of Numbers of cerata per row or arch Total living animal Anterior Posterior number of animal (mm) liver liver cerata 3 right 2,2,3 3,3,3,2,1,1 39 left 2,2,3 3,2,3,2,1,1 8 right 2,3,5,3 9,8,7,4,4 89 left 2,4,5,4 8,7,7,4,3 12 right 3,7,8,7 15,13,14,6,5,5,2 165 left 4,7,7,6 14,13,11,8,4,4,2 16 right 2,5,7,8,8 14,13,13,8,7,5,4,3,2 199 left 2,3,6,8,11 14,13,11,8,8,6,4,4,2

The jaws are almost as broad as long, and have only a very slight dorsal indentation (Fig. 9, A). There are up to 35 small teeth in a single row on the masticatory process. Two animals 8 mm in body length have 13 and 18 teeth to the radula, and a 12-mm animal has 22 teeth. The median cusp is not at all prominent as it often is in the Facelinidae, and it is flanked by four or five denticles on either side (Fig. 9, B and C). Usually the median cusp and some of the lateral denticles are finely serrate toward their bases. The anus opens beneath the first arch of cerata from the posterior liver, in the cleioproct position (Fig. 8, C). The reproductive system has several features not found in typical facelinids. The gonopore lies just behind the cerata from the anterior liver in the interhepatic space. The gonads consist of a dozen or more testis follicles each with several ovarian pouches. The hermaphrodite 1964] Edmunds: Eolid Mollusca 15

FIGURE 9. Palisa papil/ata, n. sp. A, Jaw.-B, 13th radular tooth from animal with 14 teeth.-C, Last radular tooth from animal with 22 teeth. duct runs forward to the ampulla, which lies dorsal to the rest of the genital system. Soon after leaving the ampulla, the spermoviduct divides, and the vas deferens runs through a long prostate before entering the penis. The animal from which Fig. 10 A was compiled was less than 8 mm long in life, and the prostate and penis are not of great length. Another animal 8 mm in length has an identical reproductive system except that the organs are larger; the prostate is 1 mm long and the retracted penis is '0.9 mm. When protruded by the living animal, the penis reaches a length of 1.5 mm and is very mobile; it is unarmed, and there is no accessory gland. Immediately after separating from the vas deferens, the oviduct swells into a coiled tube 0.8 mm long which is packed with spermatozoa. This presumably is the spermatheca, although in most eolids this organ is a blind pouch and not a part of the oviduct. The spermatozoa are con- centrated in the proximal part of the spermatheca, so that the part nearest the female gland mass contains only a few. At its distal end, the spermatheca divides into the oviduct and the vagina. The oviduct passes through the mucous and albuminous parts of the female genital tract, and then rejoins the vagina to form the female genital atrium. Discussion.-From the mode of branching of the liver, Pali.~a papillata is clearly a facelinid and not a favorinid (Macnae, 1954; Marcus, 1958). The Facelinidae with an unarmed penis are listed by Marcus (1958), and these genera must be compared with the present species. Caloria Trinchese has a strongly indented jaw; Facelinopsis Pruvot-Fol has an accessory penial organ; Hermissenda Bergh has the radula cusp with serrulations on its under surface; Learchis Bergh has a radula with a prominent cusp; Moridilla Bergh has cerata in rows and only two lateral dentic1es to the radula; and Rolandia Pruvot-Fol has the anus far back 16 Bulletin of Marine Science of the Gulf and Caribbean [14(1 )

h.d

rh.d B FIGURE 10. A, Pa/isa papilla/a, n. sp.-B, Phidiana Iynceus Bergh. Diagrams of reproductive system. on the right side of the animal (Bergh, 1888; Pruvot-Fol, 1954; Marcus, 1958, 1961; Marcus and Marcus, 1960). The present species differs from all existing genera of the Facelinidae in its reproductive system, particularly in the position of the spermatheca and in the vas deferens which is not prostatic in the penis. I therefore propose to make it the type species of the new genus Palisa. Phidiana lynceus Bergh Figs. 10, B; 11, A Occurrence.-This species was the commonest eolid in the mangroves at Port Royal in 1961-1962, and 88 specimens were collected. Further distribution.-St. Thomas and 81. John, Virgin Islands; Guade- loupe; Cura~ao; Brazil (Bergh, 1873, 1879; Engel, 1925; Marcus, 1955. 1957; Marcus and Marcus, 1962). Description.-The color of the living animals is similar to that described 1964] Edmunds: Eolid Mollusca 17 by Marcus (1957). The body is silvery-grey as in most eolids, but the head is suffused with orange. A line of bright vermilion-red joins the two oral tentacles, runs up their anterior margin to half of their length, and ends in a large blotch (Fig. 11, A). There are crimson markings round the mouth, and the buccal mass and the oesophagus are pale

1.0 rmt

." .',.';~~; ••• 0 0." ' . . ~'~.. l ..!~.~1~~!i>" .-~-;..!...•__:.~-~.-•• 8' .,.:::.~_...~":~' __..~-••.

((11 J: P g FIGURE 11. A, Phidiana lynceus Bergh: dorsal view with cerata removed to show liver ducts; fine stippling on head and tail indicates white, coarse stippling indicates red.-B, Learchis poica Marcus & Marcus: lateral view of living animal with cerata removed; white markings are finely stippled, yellow and orange markings coarsely stippled. crimson. There is a mid-dorsal line of white which runs from the tail to the rhinophores and bifurcates, one branch running up each of the oral tentacles. There is also a white line along the flanks. The rhinophores have a smooth, colorless, basal stalk, but the distal two-thirds of their length is irregularly annulate and has three bands of color. The annulations are rarely complete anteriorly, and they meet behind in a posterior groove. The tips of both rhinophores and oral tentacles are yellowish brown, but there are also many white flecks, and there is a white line running down the distal third of each rhinophore on the anterior face. Below this yellowish brown band on the rhinophores is a dark brown band, and the basal annulations are bright orange. The cerata often have the epidermis suffused with orange toward the tip, and there are usually two bands of white in the region of the cnidosac. The liver is nodular and basically yellow-brown, but as the nodules are blackish brown, the general appearance is dark. The ratio of tail-length to body-length is thought by Engel (1925) to be a good specific character, but Marcus (1957) shows that it varies 18 Bulletin of Marine Science of the Gulf and Caribbean [14(1) with the size of the animal. It also varies with the state of regeneration of the tail. As Marcus noticed (1957), these animals are cannibalistic, and in aquaria they frequently browse on cerata of smaller individuals. During this process, the tail is often injured, and a high proportion of animals have tails which are broken or regenerating. However, in fixed animals with apparently intact tails, the ratio of tail length to body length varies from one third to one eighth. This ratio is not, therefore, a good taxonomic character. The shape of the foot and the disposition of the cerata are similar to the description given by Marcus (1957). The posterior liver gives off several large ducts on either side, and each of these ducts supplies a number of rows of cerata (Fig. 11, A). Seven animals were dissected in order to detect the position of the anus with relation to these liver ducts. There are usually four or five raws of cerata supplied by the first duct from the posterior liver, and in six animals the anus was between the third and the fourth row of cerata (Fig. 11, A). In one animal the anus was between the fourth and the fifth row. The jaws and radular teeth closely resemble those described by Bergh (1873, 1879), Engel (1925) and Marcus (1955). The reproductive system is similar to that described by Marcus (1957), and was examined by dissection and by means of serial sections (Fig 10, B). There is no pigment in the neurilemma of the present material. Discussion.-Marcus and Marcus (1962) describe a specimen of Phidiana lynceus which was collected only 3.3 kilometers from the type locality in the Virgin Islands. They show that it is conspecific with P. selencae Bergh, and they transfer all Brazilian selencae to lynceus, which is the older name. P. brevicauda Engel differs from lynceus in that it has a shorter tail relative to its body (Engel, 1925). However, the ratio of tail length to body length is very variable in the present material, even in animals which have undamaged tails, and this cannot therefore be used as a taxonomic character. I therefore consider brevicauda Engel together with selencae Bergh to be synonyms of Phidiana lynceus Bergh.

Learchis poica Marcus and Marcus Fig. 11, B

Occurrence.-Twelve specimens were found in October 1961 and In February and March 1962 at Port Royal. Further distribution.-Miami, Florida (Marcus and Marcus, 1960). Description.-The living animal was not described by Marcus and Marcus (1960), and it is therefore described here for the first time (Fig. 11, B). 1964] Edmunds: Eolid Mollusca 19 The length of the living animal is up to 12 mm; the following description was taken from an animal 9 mm long. The oral tentacles and rhino- phares both reach 2.5 mm, the cerata reach 3 mm, and the angles of the foot, which is not notched, are 0.5 mm. The tail is 1.5 to 2 mm long. The general body color is silvery-grey. There is a white patch on the forehead and another just behind the rhinophores. There is a mid- dorsal line and lateral lines of irregular white markings. There is an orange cheek patch and often a pale orange line from the rhinophorcs to the oral tentacles. In some animals there is a lateral line of yellow in the angle of the foot with the flanks. The rhinophores have irregular rings some of which are incomplete posteriorly, and some resemble tubercles rather than the annulations of typical facelinids. The bases of both rhinophores and oral tentacles are tinged with yellowish. The tips are covered with white blotches. The tail is spotted with white, and there may be white flecks elsewhere on the body and on the angles of the foot. The cerata are covered with white blotches, and there is a white ring in the cnidosac region. The liver is creamy yellow, with brownish red toward the tip, at the base, and as small spots in between. The renal pore may be orange or crimson, and the gonopore may have a pale orange rim. The distribution of cerata and the positions of the anal, renal and genital apertures is the same as in the description given by Marcus and Marcus (1960). The jaws and radula were examined from two animals, and closely resemble the figures given by Marcus and Marcus. The median cusp of the radular tooth is very long and has from 4 to 9 denticles on each side. In one of the animals, these denticles are very irregular in shape and size. Such irregularities occur occasionally in other eolid species, but the cause is not known. The reproductive system resembles that of the original material (Marcus and Marcus, 1960), the most important features being the penial gland opening separately from the prostatic vas deferens, the unarmed penis, and the spermatheca with a folded wall opening close to the junction of the vas deferens with the oviduct. The larva has a spiral shell of Thompson's type 1, as in Facelina auriculata (Thompson, 1961).

Family Favorinidae Subfamily Favorininae Favorinus auritulus Marcus Figs. 12; 13; 14, A Occurrence.-Five specimens were taken at Port Royal in 1961-62, and a single animal was found at Miami in 1962. Further distribution.-Brazil (Marcus, 1955). 20 Bulletin of Marine Science of the Gulf and Caribbean [14(1) Description.- The living animals are up to 8 mm long, with the cerata reaching 3 mm, the rhinophores and the oral tentacles 2.5 mm, the foot angles 1 mm, and the tail 3 mm. The body is very transparent and the shape of the cerebral ganglia is clearly visible in the living animal. The base of the rhinophores is brown or yellowish brown, as described by Marcus (1955), and the tips of rhinophores, oral and foot tentacles are white. There are additional white spots on the body and cerata, the tail is white, and there may be one or two white bands at the ceras tips. The liver may be pink or creamy yellow with brown or reddish brown spots. The general appearance of the liver thus varies from pinkish or dirty yellow to dark brown. The liver ducts in the body are clearly visible in the living animal. The cnidosac is white, and its connecting duct to the liver is usually dark brown (Fig. 12, C).

I'X~ H

1.0 :::::1 .' .-, '\--g'------' " ell an *

FIGURE 12. Favorinus auritulus Marcus. A, Dorsal view of living animal with right cerata removed. B, Ventral view of head to show shape of foot. C, Ceras: white markings are stippled; the liver is creamy or pinkish with reddish brown spots, and a dark spot just below the cnidosac.

There are two bulbs on each of the rhinophores, although in young animals these are very inconspicuous (Fig 12, A). According to Marcus (1955), the bulbs may be absent in young specimens, but develop with increasing size. The foot is rounded anteriorly, without a notch, and lacks a marked'longitudinal groove (Fig. 12, B). The cerata are arranged as described by Marcus (1955). There are 31 in an animal 2.5 mm in length, and up to 78 in animals 8 mm long. (Fig. 12, A). The jaws (Fig. 13, A) resemble those described by Marcus (1955) but are less broad. There are three or four rows of denticles (Fig. 13, B), 1964] Edmunds: Eolid Mollusca 21

FIGURE 13. Favorinus auritulus Marcus. A, Jaw.-B, DenticIes on masticatory border of jaw.-C, Surface view of 9th radular tooth of animal with 18 teeth.-D, Side view of 15th tooth from same animal.

and the total number of the largest denticles is 82. There are 18 teeth in the radula excluding one "ghost" tooth in process of formation. The radular teeth have no lateral denticles as have those reported by Marcus (1955) (Fig. 13, C-D). The reproductive system was examined from serial sections (Fig. 14, A). There are a dozen or more ovotestis sacs which collect into a median hermaphrodite duct. This swells into the 0.6 mm-Iong tubular ampulla, and the spermoviduct divides soon after leaving the ampulla into the vas deferens and the oviduct. The vas deferens is prostatic for most of its length (0.5 mm), and it opens at the tip of the unarmed penis. The penis in one animal is 100fL long and 75fL broad, whilst in another animal it is 190fL long and 95fL broad. The oviduct gives off a spermatheca 0.2 mm in diameter just before it enters the glandular part of its length. Discussion.-The Favorininae have only one row of cerata to each liver branch (Marcus, 1958). Favorinus (luritulus is very similar to the European F. branchialis (Rathke), but differs in the shape of the penis (Marcus, 1955). According to Marcus, the penis is five times as long as it is broad in auritulus, whilst the length and breadth are the same in branchialis. In the present material the ratio of length to breadth of the penis is 4/3 in one animal and 2 in another. It is therefore intermediate between the two ratios given by Marcus, and it is likely that this may prove not to be a good taxonomic character. The Jamaican animals differ from the Brazilian F. auritulus in two other features: the presence 22 Bulletin oj Marine Science oj the Gulj and Caribbean [14(1)

h'dl

100 •• h'l '------' ~O('O A It B FIGURE 14. A, Favorinus auritulus Marcus.-B, Godiva mbrolineata, n. sp. Diagrams of reproductive system. of two rather than three bulbs to the rhinophores; and the absence of lateral denticles on the radular teeth. The number of rhinophorial bulbs is probably not important since it may indicate that the Jamaican animals were not fully grown. Also, there is considerable variation in F. branchialis with respect to shape and the presence or absence of rhinophorial bulbs (Labbe, 1929). Normally, this species has only one bulb, but, together with typical animals, I found one specimen which had three well-marked bulbs at Millport, Scotland, in 1959. If this degree of variation can occur in one species of Favorinus, then there is no reason it should not also occur in another species. In the case of the radular teeth, Labbe (1929) described several well-marked varieties of F. branchialis, some of which have lateral denticles to the radular teeth and some of which lack them. According to Labbe and to Alder and Hancock (1845-1855), the British variety has no lateral denticles, but I have a specimen collected at Plymouth in 1960 which does possess 1964] Edmunds: Eolid Mollusca 23 lateral denticles. Since F. branchialis mayor may not have lateral denticles on the radular teeth, it is likely that a similar degree of variation may occur in a closely related species. There can thus be little doubt that the Jamaican material is Favorinus auritulus. It should be stressed, however, that F. auritulus and branchialis are very closely related. They differ chiefly in size and color. F. branchialis frequently exceeds 20 mm alive, whilst F. auritulus has not been found longer than 12 mm. Small branchialis often approach auritulus in color, although the larger animals are quite distinct. It is probable that if auritulus had been found on the east coast of the Atlantic instead of the west, it would have been regarded as a variety of F. branchialis, and not as a distinct species. It is possible that F. branchialis occurs on both sides of the Atlantic, since at least two species of eolid have a trans-Atlantic distribution (Spurilla neapolitana and Berghia coerulescens), but until more material is available, J prefer to regard auritulus and branchialis as separate species. More data are wanted on the proportions of the penis, and on the size and markings of the living animal. It would be interesting, too, to know whether F. auritulus feeds on eggs of other opisthobranchs as is reported for F. branchialis (Alder and Hancock, 1845-1855; Haefelfinger, 1962).

Subfamily Facalaninae Godiva rubrolineata, n. sp. Figs. 14, B; 15; 16 OcclIrrence.-One animal was taken at Port Royal in December 1961; seven more, including the holotype, were collected at Miami in April 1962. The holotype, fixed in Bouin's fluid, is deposited in the British Museum (N.H.), no. 1963117 W. Description.- The living animals are up to 9 mm long, with the oral tentacles 2.5 mm, rhino phares 2.3 mm, foot angles 1.3 mm, cerata 1.5 mm, and tail 3.5 mm. The rhinophores are smooth but may wrinkle slightly when they are contracted (Fig. 15, A). The foot is notched and longitudinally grooved (Fig. 15, B). The general body color is silvery gray. There is white coloration dorsally on the tail, the foot angles, the upper two-thirds of the rhinophores, and the upper half of the oral tentacles. The Florida material also has scattered white dots on the head and body. There is a bright orange-red line which runs from in front of the rhinophores down the back of the animal. It is broken at the levels of the ceras groups (Fig. 15, A). There is also an orange- red cheek marking, and in most cases this continues laterally along the flanks as a broken line similar to the dorsal stripe (Fig. 15, C). The cerata are pointed (Fig. 15, D), with a white cnidosac, and with 24 Bulletin of Marine Science of the Gulf and Caribbean [14(1)

1.0 alii 0.5 tJITl '------'

r. p

FIGURE 15. Godiva rubrolineata, n. sp. A, Dorsal view of living animal; white markings finely stippled, orange-red markings coarsely stippled.-B, Ventral view of head to show notched foot.-C, Lateral view with right cerata removed; stippling as in A.-D, Ceras, with blackish liver, white cnidosac, and scattered white spots. scattered white dots in the Jamaican animal, but with an anterior white line in some of the Florida animals. The liver is basically buffish, but this is largely obscured by black in the Jamaican, and by brown or red-brown in the Florida animals. The cerata from the anterior liver form a two-rowed arch, as in other species of the Pacalaninae (Marcus, 1958). There are four or five branches to the posterior liver on each side; the front ones are in single or double arches, the rear ones are in simple rows (Fig. 15, C). The cerata are distributed as follows: Anterior liver Posterior liver Jamaican specimen: Right side 12 10, 6, 4, 2 Left side 12 8, 3, 4, 1 Florida specimen: Right side 11 8, 5, 3, 3, 1 Left side 10 7, 5, 5, 2, 1 The anus in all animals opens under the first arch of the right posterior 1964] Edmunds: EoUd Mollusca 25 liver; the renal pore is interhepatic; the gonopore is below the cerata from the anterior liver (Fig. 15, C). The jaws have a small dorsal indentation (Fig. 16, A), and in three animals examined have from 23 to 29 denticles in a single row on the masticatory border (Fig. 16, B). The same three animals have from 21 to 26 teeth in the radula. The median cusp is strong and is flanked by four or five long, curved denticles (Fig. 16, C). In the Jamaican animal these denticles are slightly thicker than are those from the Florida animals.

FIGURE 16. Godiva rubrolineata, n. sp. A, Jaw.-B, Denticles on masticatory border of jaw.-C, 18th tooth of an animal with 21 radular teeth.

The reproductive system was examined from serial sections of three animals. Figure 14 B is taken from an animal in the male phase; hence it lacks well-developed eggs in the ovotestes, there are few spermatozoa in the spermatheca, and the female gland mass is poorly developed. In this animal there are 8 ovotestis follicles which open into the hermaphrodite duct. The ampulla is slightly coiled and 3501L long. Soon after leaving the ampulla, the spermoviduct divides. The vas deferens is prostatic for most of its length, and runs for 0.4 mm before entering the penis, which is 0.1 mm long. The latter opens obliquely at its unarmed tip. The oviduct receives a duct from the 1501L-Iongspermatheca before it enters the female gland mass. The male and female gonopores are separated by a thin fold. In a larger animal, in which the ovaries were functional, the genital system is very similar, but some of the organs are larger. Thus, the ampulla reaches 0.6 mm, the spermatheca 0.45 mm, the prostate 0.55 mm, the penis 0.4 mm, and there are 11 ovotestis follicles. Discussion.-The classification of the Facalaninae is in an unsatisfactory state. Apart from Falcalana Bergh, which can be distinguished by the shape of its mandibles, the remaining four genera are all diagnosed on the nature of the penial armature. It is clear from the descriptions of the type-species of these genera that the male genital system is subject to 26 Bulletin of Marine Science of the Gulf and Caribbean [14(1) considerable variation in this subfamily, and that the armature of the penis is only one of several characters. Equally important taxonomic characters are the presence or absence of a penial gland and the nature of the prostate. In Godiva quadricolor (Barnard) the vas deferens is prostatic (Macnae, 1954); in Dondice banyulensis Portmann and Sand- meier the vas deferens has two coiled regions which appear to be glandular from the illustration (Portmann and Sandmeier, 196'0); and in Dondice occidentalis (Engel) the vas deferens is not prostatic, but the prostate is present as a separate pouch (Marcus, 1958). The species of the Facalaninae in which the male genital system is at all well known are listed in Table 2. The type-species of the genera Dondice Marcus, Godiva Macnae, Austraeolis Burn and Echinopsole Macnae are given first, and the differences between them can be seen clearly.

TABLE 2 PENIAL CHARACTERSOF SPECIES OF FACALANINAE For further explanation, see text. Penial Penial Armature Gland Prostate Gland Dondice occidentalis (Engel) none present as a separate pouch Godiva quadricolor (Barnard) a hook absent part of vas deferens Austraeolis ornata (Angas) fleshy unknown unknown filaments Echinopsole fulvus Macnae many present unknown spines Echinopsole breviceratae many present unknown Burn spines Godiva rubrolineata, n. sp. none absent part of vas deferens Dondice banyulensis Portmann & Sandmeier none absent part of vas def. (2 parts) Dondice modesta (Bergh) none present? unknown Favorinus horridus Macnae none absent part of vas deferens

It is evident that until the male genital systems of more species are known, it is difficult to decide which characters should be given generic and which should be given specific rank. Nevertheless, we can draw some conclusions on the basis of the species at present known. The Australian species breviceratae Burn closely resembles Echinopsole fulvus Macnae, and Burn (1962) correctly assigns it to this genus. 1964] Edmunds: Eolid Mollusca 27 The Caribbean rubrolineata differs from Dondice occidenta/is in two out of three characters, the penial gland and the prostate, but it differs from Godiva quadricolor in only one out of three characters, the penial armature (Marcus, 1958; Macnae, 1954). I therefore allocate it to the genus Godiva. Macnae's original diagnosis of Godiva (Macnae, 1954) includes species with or without a penial hook, and with this I am in agreement. But the diagnosis should now be extended to include the absence of a penial gland and the presence of a prostatic vas deferens. The recently described Mediterranean species banyulensis Portmann and Sandmeier is placed in the genus Dondice (Portmann and Sandmeier, 1960), but Dondice is not the only genus with an unarmed penis. Dondice must also have a penial gland, and perhaps also the prostate as a separate pouch. The species banyulensis possesses neither of these features, and it thus resembles the genus Godiva. It differs from G. quadricolor and G. rubrolineata in that the vas deferens has two coiled regions instead of one, but this could be merely a specific difference. Although it is not known if either of these regions is prostatic, it is therefore probable that banyulensis Portmann and Sandmeier belongs to the genus Godiva. Favorinus horridus Macnae is transferred to the Facalaninae by Marcus (1958) because it has multiserial rows of cerata. As can be seen from Table 2, it clearly belongs to the genus Godiva. When more species of Godiva with an armed penis are known, the three species discussed here, rubrolineata, banyulensis and horridus, may be excluded from Godiva; but at present I consider this to be unnecessary. Rizzolia modesta Bergh is placed in the genus Dondice by Marcus (1958). The nature of the prostate in this species is not known, but there is a stalked body near the penis which could be either a penial gland or a prostate pouch (Bergh, 1881). If this proves to be correct. then modesta Bergh does indeed belong to Dondice.

Dondice occidentalis (Engel) Occurrence.-ll specimens were taken between November 1961 and March 1962 at Port Royal. Further distribution.-Montego Bay, Jamaica (Engel, 1925); Brazil (Marcus, 1958); Florida (Marcus and Marcus, 1960). Description.-The present animals agree closely with the description given by Marcus (1958), but there are several small differences. In the living animal there is considerable color variation, but the most important characters seem to be the mid-dorsal stripe of orange-red from the level of the eyes forward, and the lateral stripes of the same color running from the base of the oral tentacles back to the tail. Sometimes these stripes are orange or even brownish yellow, but they are present in both Jamaican and 28 Bulletin of Marine Science of the Gulf and Caribbean [14(1) Brazilian animals (Marcus, 1958). The body may be suffused with brown, yellowish brown or pinkish brown, and the back may bear scattered minute black flecks. There may be a mid-dorsal white line which may be broken into a series of spots. There may be a white lateral line at the level of the ceras insertions, or this may be absent, and there are usually white spots scattered over the surface of the body. The Jamaican animals lack the orange-red ring on the cerata that Marcus describes (1958). The liver in the cerata is usually brownish yellow with darker markings, and the ceratal epidermis may have white bands or white blotches. The location of the gonopores differs slightly from that in Brazilian animals. Marcus (1958) describes the genital apertures as lying under the arch of cerata from the anterior liver, but in the present animals they are located immediately behind this arch, at the front of the interhepatic space. The orange-red stripe passes ventral to the genital apertures. Another difference from the Brazilian animals is that in life the Jamaican animals have no foot notch. The jaws are covered with a black epithelium exactly as Marcus describes (1958), so it is probable that Engel's material from Jamaica also has this character although he does not mention it in his description (Engel, 1925). The jaw has a conspicuous dorsal indentation, and both jaw and radula agree with the descriptions given by Marcus and Engel. The reproductive system is very similar to that of the Brazilian animals (Marcus, 1958). The important characters are the prostate which opens independently from the vas deferens, and the presence of an accessory male, or penial, gland opening into the vas deferens. In the present material it opens into the vas deferens at the base of the penis, and the vas deferens continues to be dandular almost to its aperture. In Brazilian animals the junction of the accessory gland and the vas deferens is nearer the tip of the penis. Discussion.-It is possible that these differences between the Jamaican and the Brazilian animals warrant specific or subspecific separation, but until specimens from elsewhere in the Caribbean are available, I am inclined to follow Marcus (1958) and call both animals Dondice occi- c'entalis (Engel). Family Aeolidiidae Spurilla neapolitana (Delle Chiaje) Occurrence.-Seventeen animals were found at Port Royal where this species is the commonest of the anemone-feeding eolids. Two more were found at Miami. Further distribution.-Mediterranean; Atlantic coast of south Europe; Brazil; West Indies; Florida (Engel, 1925; Pruvot-Fol, 1954; Marcus, 1955; Marcus and Marcus, 1960). 1964] Edmunds: Eolid Mollusca 29 Discussion.-The present animals vary in size from 3.5 to 37 mm in life. They are easily recognized by the large circular white spots on the body and cerata, the orange tinge to the epidermis and the viscera, and the usually greenish liver diverticula in the cerata. The radula and jaws were examined to confirm the identification. Following Marcus and Marcus (l960), the American animals are no longer separated from the European ones as var. braziliana. Berghia eoeruleseens (Laurillard) Oeeurrenee.-Three animals were found at Port Royal, and one was found at Miami. Further distribution.-Mediterranean; Atlantic coast of south Europe; Tobago, West Indies; Brazil (Engel, 1925; Pruvot-Fol, 1954; Marcus, 1957). Diseussion.- This is a very variable species, but there is usually orange on the head and at the ceras insertions. The ring in the cnidosac region may be white, yellow or orange, and the liver may be greenish or dark brown. The largest animal was 40 mm long in life. The radula and jaws were examined to confirm the identification. A recent description of this species is given by Marcus, 1957. ZOOGEOGRAPHY Of the thirteen species of eolid here described from Jamaica, ten are known from Miami, Florida, and eight from the coast of Sao Paulo, Brazil. This supports the inclusion of the tip of Florida and the southern coast of Brazil in the same faunistic area as the Caribbean (Hedgpeth, 1957; Marcus and Marcus, 1960). None of the present species is known from the Atlantic coast of America north of Florida, but two species are trans-Atlantic in their distribution. ECOLOGY Seventeen collections were made from the mangrove channel at Port Royal during a period of five and one-half months, but there is little evidence that any of the species are seasonal in their occurrence. The common species, after their initial discovery in October or November 1961, were found more or less regularly until collections ceased in March 1962. The exceptions are Catdona maua, C. perea and Learehis poica. In the case of C. maua, 20 animals were found in two collections on October 10th and 17th, and then none were found until February 22nd. Between then and March 21st, 12 more animals were found in four collections. L. poica was similarly absent from 17th October until 22nd February. This distribution could indicate that the two species are cyclic in their occurrence, but there is a more likely explanation. Goodbodv (1961) describes the effect of heavy tropical rains on the fauna of the mangrove channel in which these eolids were found. There were similar 30 Bulletin of Marine Science of the Gulf and Caribbean [14(1) heavy rains in late October 1961 (Edmunds, 1963), and it is probable that the resultant reduction in salinity caused mortality of many inverte- brates in the area. It is likely that this killed off both Catriona maua and Learehis poiea. The case of C. perea is different, since 31 specimens were found in five colIections between November 23rd and December 19th, but none was found in the folIowing three months. There is thus some evidence that Catriona perea is cyclic in its occurrence in Jamaica. Out of this large collection of eolids, four individuals were found to be parasitized by pycnogonids. One pycnogonid was found on Catriona perea, and the other three were found on Spurilla neapolitana. In all four cases, the proboscis of the pycnogonid was inserted into the back or the side of the eolid, and in one case it was definitely inserted into the liver duct on the back of a large S. neapolitana. Since pycnogonids feed by sucking soft-bodied animals, including hydroids, sponges, and even bivalved molluscs (Ohshima, 1937; Fage, in Grasse, 1949), it is highly probable that they were feeding on the eolids. ACKNOWLEDGMENTS I should like to thank Professor D. M. Steven for providing facilities for colIecting and studying material in the Department of Zoology of the University of the West Indies, Jamaica; Dr. F. M. Bayer and Dr. H. B. Owre for helping with the colIection at Miami; and Professor J. W. S. Pringle for providing laboratory facilities at Oxford. Dr. D. Nichols, Dr. A. J. Cain and my wife Janet alI read parts of the manuscript, and I am most grateful to them. I also wish to thank the Department of Scientific and Industrial Research and the Committee for Commonwealth University Interchange for providing grants for research and travel. POSTSCRIPT Since the manuscript of this paper was submitted for publication. an article entitled "Measuring nudibranchs: a standardization for descriptive purposes" has appeared (Risso-Dominguez, C.J., 1963, Proc. malac. Soc. Lond., 35: 193-202). Using the terminology proposed therein, all measurements of the lengths of eolids described in this paper refer to the distance "Ac." SUMARIO MOLUSCO EOUDO DE JAMAICA, CON DESCRIPCIONES DE DOS NUEVOS GENEROS Y TRES NUEVAS ESPECIES Este trabajo describe la presencia de trece especies de nudibranquios e6lidos de Jamaica. La mayoria del material fue co1eccionado entr'~ Octubre de 1961 y Marzo de 1962 en mangles de Port Royal, pero algunos ejemplares fueron obtenidos en otras partes de Port Royal y tambien en Miami, Florida. Dos nuevos generos y tres nuevas especies de nudibranquios e61idos son descritos. EI nuevo genero Selva tiene un ano cleioprocto pero esta estrechamente 1964J Edmunds: Eolid Mollusca 31 relacionado con Catriona, Cuthona y Tergipes y por tanto pertenece a la acIeioprocta Cuthonidae. La especie tipo Selva rubra sp. n. es descrita basandose en tres ejemplares de Port Royal. Palisa gen. n. tiene los rin6foros tuberculados y la espermateca forma parte del oviducto. La especie tipo Palisa papillata sp. n. es descrita bas an dose en veinticuatro ejemplares procedentes de Port Royal y uno de Miami. La tercera nueva especie, Godiva rubrolineata es conocida por siete ejemplares de Miami y uno de Jamaica. EI genero Godiva Macnae, especie tipo G. quadricolor (Barnard), es revisada para incluir rubrolineata, banyulensis (Portmann y Sandmeier) y horridus (Macnae). De las trece especies de e61idos conocidos de Jamaica, diez son tambien conocidos en la Florida, ocho de Sao Paulo y dos de Europa. Hay cierta evidencia de que Catriona perea y quizas C. maua y Learchis poiea tambien son cicIicos en su ocurrencia, pero no se sabe si las apari- ciones de estas especies son anuales 0 irregulares. Cuatro casos de e6lidos parasitados por pignog6nidos son descritos, uno de C. perea y tres de Spurilla neapolitana.

REFERENCES ALDER, JOSHUA AND ALBANY HANCOCK 1845-55. A monograph of the British nudibranchiate Mollusca. Ray Society, London, [xiil+54 pp, 48 pIs. and explanatory text without pagin- ation, xl. BERGH, R. 1873. Beitrage zur Kenntnis der Aeolidiaden. 1. Verh. zool.-bot. Ges. Wien, 23: 597-628. 1879. Beitrage zur Kenntnis der Aeolidiaden. 6. Verh. zool.-bot. Ges. Wien, 28: 553-584. 1881. Beitrage zur Kenntnis der japanischen Nudibranchien. 1. Verh. zool.-bot. Ges. Wien, 30: 155-200. 1888. Beitrage zur Kenntnis der Aeolidiaden. 9. Verh. zool.-bot. Ges. Wien, 38: 673-706. BURN, R. 1962. Descriptions of Victorian nudibranchiate Mollusca, with a compre- hensive review of the Eolidacea. Mem. nat. Mus. Victoria, 25: 95- 128. EDMUNDS, MALCOLM 1963. Berthelinia caribbea n. sp., a bivalved gastropod from the west Atlantic. J. Linn. Soc. (Zoo!.), 44: 731-739. ENGEL, H. 1925. Westindische opisthobranchiate Mollusken. 1. Aeolidiadae. Bijdr. Dierk., 24: 33-80. 1927. Westindische opisthobranchiate Mollusken. 2. Aplysiidae, Pleuro- branchidae, Oxynoeidae, Elysiidae, Phyllobranchidae. Bijdr. Dierk., 25: 83-122. GOODBODY, IVAN 1961. Mass mortality of a marine fauna following tropical rains. Ecology, 42: 150-155. 32 Bulletin of Marine Science of the Gulf and Caribbean [14(1)

GRASSE, P. 1949. Traite de Zoologie, 6. Masson, Paris, 979 pp. HAEFELFINGER, H. R. 1962. Quelques faits concernant la nutrition chez Favorinus branchialis (Rathke 1806) et Stilige>rvesiculosus (Deshayes 1864), deux Mollus- ques Opisthobranches. Rev. suisse Zoo!., 69: 312-316. HECHT, E. 1896. Contribution a l'etude des nudibranches. Mem. Soc. zoo!. Fr., 8: 537-711. HEDGPETH, JOEL W. 1957. Marine biogeography. Mem. geo!. Soc. Amer., 67: 359-382. LABBE, A. 1929. Les organes palleaux (caryophyllidies) et Ie tissu conjunctif du manteau de Rostanga coccinea Forbes. Arch. Anat. micr., 25: 87-103. MACNAE, W. 1954. On some eolidacean nudibranchiate molluscs from South Africa. Ann. Natal Mus., 13: 1-50. MARCUS, ERNST 1955. Opisthobranchia from Brazi!. Bol. Fac. Filos. Cienc. S. Paulo, 20: 89-262. 1957. On Opisthobranchia from Brazil. J. Linn. Soc. (Zool.), 43: 390-486. 1958. On western Atlantic opisthobranchiate gastropods. Amer. Mus. Novi- tates, no. 1906, 82 pp. 1961. Opisthobranch mollusks from California. Veliger, 3 (Supp!.): 1-84. MARCUS, ERNST AND EVELINE MARCUS 1960. Opisthobranchs from American Atlantic warm waters. Bull Mar. Sci. Gulf & Carib., 10: 129-203. 1962. Opisthobranchs from Florida and the Virgin Islands. Bull. Mar. Sci. Gulf & Carib., 12: 450-488. ODHNER, NILS HJ. 1939. Opisthobranchiate Mollusca from the western and northern coasts of Norway. K. norske vidensk. Selsk. Skr., no. 1, 93 pp. OHSHIMA, H. 1937. The life-history of Nymphonella tapetis Ohshima (Pantopoda, Eury- cydidae). C.R. 12th Congr. Int. Zool., 1616-1626. PORTMANN, A. AND E. SANDMEIER 1960. Dondice banyulensis, sp. nov. un Eolidien nouveau de la Mediter- ranee. Rev. suisse Zoo!., 67: 159-168. PRUVOT-POL, ALICE ] 954. Mollusques Opisthobranches. Paune de France, 58. Lechevalier, Paris, 460 pp. RISBEC, JEAN 1937. Note preliminaire au sujet de nudibranches Neo-Caledoniens. Bull. Mus. Hist. nat., Paris, 9: 159-164. 1953. Mollusques nudibranches de la Nouvelle-Caledonie. Paune de l'Union Pranc;:aise, 15. 189 pp. SMALLWOOD, W. M. 1910. Notes on the hydroids and nudibranchs of Bermuda. Proc. zoo!. Soc. Lond., 137-145. THOMPSON, T. E. 1961. The importance of the larval shell in the classification of the Sacoglossa and the Acoela (Gastropoda Opisthobranchia). Proc. malac. Soc. Lond., 34: 233-238.